The last common ancestor (LCA) shared by chimpanzees (Pan troglodytes) and bonobos (P. paniscus) was an Early Pleistocene African ape, which, based on the behavior of modern chimpanzees, may be assumed to be a tool-using animal. However, the character of tool use in the Pan lineage prior to the 20th century is largely unknown. Here, I use available data on wild bonobo tool use and emerging molecular estimates of demography during Pan evolution to hypothesise the plausible tool use behavior of the bonobo-chimpanzee LCA (or "Pancestor") at the start of the Pleistocene, over 2 million years ago. This method indicates that the common ancestor of living Pan apes likely used plant tools for probing, sponging, and display, but it did not use stone tools. Instead, stone tool use appears to have been independently invented by Western African chimpanzees (P. t. verus) during the Middle Pleistocene in the region of modern Liberia-Ivory Coast-Guinea, possibly as recently as 200,000-150,000 years ago. If this is the case, then the LCA of humans and chimpanzees likely also did not use stone tools, and this trait probably first emerged among hominins in Pliocene East Africa. This review also suggests that the consistently higher population sizes of Central African chimpanzees (P. t. troglodytes) over the past million years may have contributed to the increased complexity of wild tool use seen in this sub-species today.

We recorded the damage that wild bearded capuchin monkeys (Sapajus libidinosus) caused to a sandstone anvil during pounding stone tool use, in an experimental setting. The anvil was undamaged when set up at the Fazenda Boa Vista (FBV) field laboratory in Piauí, Brazil, and subsequently the monkeys indirectly created a series of pits and destroyed the anvil surface by cracking palm nuts on it. We measured the size and rate of pit formation, and recorded when adult and immature monkeys removed loose material from the anvil surface. We found that new pits were formed with approximately every 10 nuts cracked, (corresponding to an average of 38 strikes with a stone tool), and that adult males were the primary initiators of new pit positions on the anvil. Whole nuts were preferentially placed within pits for cracking, and partially-broken nuts outside the established pits. Visible anvil damage was rapid, occurring within a day of the anvil's introduction to the field laboratory. Destruction of the anvil through use has continued for three years since the experiment, resulting in both a pitted surface and a surrounding archaeological debris field that replicate features seen at natural FBV anvils.

Animals in captive or laboratory settings may outperform wild animals of the same species in both frequency and diversity of tool use, a phenomenon here termed captivity bias. Although speculative at this stage, a logical conclusion from this concept is that animals whose tool-use behaviour is observed solely under natural conditions may be judged cognitively or physically inferior than if they had also been tested or observed under controlled captive conditions. In turn, this situation creates a potential problem for studies of the behaviour of extinct members of the human family tree-the hominins-as hominin cognitive abilities are often judged on material evidence of tool-use behaviour left in the archaeological record. In this review, potential factors contributing to captivity bias in primates (including increased contact between individuals engaged in tool use, guidance or shaping of tool-use behaviour by other tool-users and increased free time and energy) are identified and assessed for their possible effects on the behaviour of the Late Pleistocene hominin Homo floresiensis. The captivity bias concept provides one way to uncouple hominin tool use from cognition, by considering hominins as subject to the same adaptive influences as other tool-using animals.

Tool use is a vital component of the human behavioural repertoire. The benefits of tool use have often been assumed to be self-evident: by extending control over our environment, we have increased energetic returns and buffered ourselves from potentially harmful influences. In recent decades, however, the study of tool use in both humans and non-human animals has expanded the way we think about the role of tools in the natural world. This Theme Issue is aimed at bringing together this developing body of knowledge, gathered across multiple species and from multiple research perspectives, to chart the wider evolutionary context of this phylogenetically rare behaviour.

Burmese long-tailed macaques (Macaca fascicularis aurea) are one of a limited number of wild animal species to use stone tools, with their tool use focused on pounding shelled marine invertebrates foraged from intertidal habitats. These monkeys exhibit two main styles of tool use: axe hammering of oysters, and pound hammering of unattached encased foods. In this study, we examined macroscopic use-wear patterns on a sample of 60 wild macaque stone tools from Piak Nam Yai Island, Thailand, that had been collected following behavioural observation, in order to (i) quantify the wear patterns in terms of the types and distribution of use-damage on the stones, and (ii) develop a Use-Action Index (UAI) to differentiate axe hammers from pound hammers by wear patterns alone. We used the intensity of crushing damage on differing surface zones of the stones, as well as stone weight, to produce a UAI that had 92% concordance when compared to how the stones had been used by macaques, as observed independently prior to collection. Our study is the first to demonstrate that quantitative archaeological use-wear techniques can accurately reconstruct the behavioural histories of non-human primate stone tools.

The dispersal of Homo sapiens out of Africa is a significant topic in human evolutionary studies. Most investigators agree that our species arose in Africa and subsequently spread out to occupy much of Eurasia. Researchers have argued that populations expanded along the Indian Ocean rim at ca 60,000 years ago during a single rapid dispersal event, probably employing a coastal route towards Australasia. Archaeologists have been relatively silent about the movement and expansion of human populations in terrestrial environments along the Indian Ocean rim, although it is clear that Homo sapiens reached Australia by ca 45,000 years ago. Here, we synthesize and document current genetic and archaeological evidence from two major landmasses, the Arabian peninsula and the Indian subcontinent, regions that have been underplayed in the story of out of Africa dispersals. We suggest that modern humans were present in Arabia and South Asia earlier than currently believed, and probably coincident with the presence of Homo sapiens in the Levant between ca 130 and 70,000 years ago. We show that climatic and environmental fluctuations during the Late Pleistocene would have had significant demographic effects on Arabian and South Asian populations, though indigenous populations would have responded in different ways. Based on a review of the current genetic, archaeological and environmental data, we indicate that demographic patterns in Arabia and South Asia are more interesting and complex than surmised to date.

Genetic studies of South Asias population history have led to postulations of a significant and early population expansion in the subcontinent, dating to sometime in the Late Pleistocene. We evaluate this argument, based on new mtDNA analyses, and find evidence for significant demographic transition in the subcontinent, dating to 35-28 ka. We then examine the paleoenvironmental and, particularly, archaeological records for this time period and note that this putative demographic event coincides with a period of ecological and technological change in South Asia. We document the development of a new diminutive stone blade (microlithic) technology beginning at 35-30 ka, the first time that the precocity of this transition has been recognized across the subcontinent. We argue that the transition to microlithic technology may relate to changes in subsistence practices, as increasingly large and probably fragmented populations exploited resources in contracting favorable ecological zones just before the onset of full glacial conditions.

All modern humans use tools to overcome limitations of our anatomy and to make difficult tasks easier. However, if tool use is such an advantage, we may ask why it is not evolved to the same degree in other species. To answer this question, we need to bring a long-term perspective to the material record of other members of our own order, the Primates.

We present a superalgebraically convergent integral equation algorithm for evaluation of TE and TM electromagnetic scattering by smooth perfectly conducting periodic surfaces z=f(x). For grating-diffraction problems in the resonance regime (heights and periods up to a few wavelengths) the proposed algorithm produces solutions with full double-precision accuracy in single-processor computing times of the order of a few seconds. The algorithm can also produce, in reasonable computing times, highly accurate solutions for very challenging problems, such as (a) a problem of diffraction by a grating for which the peak-to-trough distance equals 40 times its period that, in turn, equals 20 times the wavelength; and (b) a high-frequency problem with very small incidence, up to 0.01 degrees from glancing. The algorithm is based on the concurrent use of Floquet and Chebyshev expansions together with certain integration weights that are computed accurately by means of an asymptotic expansion as the number of integration points tends to infinity.

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